Drift is often characterized in statistical terms. Yet such a purely statistical characterization is ambiguous for it can accept multiple physical interpretations. Because of this ambiguity it is important to distinguish what sorts of processes can lead to this statistical phenomenon. After presenting a physical interpretation of drift originating from the most popular interpretation of fitness, namely the propensity interpretation, I propose a different one starting from an analysis of the concept of drift made by Godfrey-Smith. Further on, I show (...) how my interpretation relates to previous attempts to make sense of the notion of expected value in deterministic setups. The upshot of my analysis is a physical conception of drift that is compatible with both a deterministic and indeterministic world. (shrink)

Building upon a non-standard understanding of evolutionary process focusing on variation and persistence, I will argue that communities and ecosystems can evolve by natural selection as emergent individuals. Evolutionary biology has relied ever increasingly on the modeling of population dynamics. Most have taken for granted that we all agree on what is a population. Recent work has reexamined this perceived consensus. I will argue that there are good reasons to restrict the term “population” to collections of monophyletically related replicators and (...) interactors, which explains why many existing models in population biology exclude by definition many genuine evolving biological individuals such as communities and ecosystems. By studying a case of community evolution (a symbiotic termite–fungus community), we will see that it is variation that is important to evolutionary processes, not populations. Variation within a population is only one of many types of variation that can lead to evolution by natural selection. The upshot of focusing on variation is that cases of community and ecosystem adaptive change become tractable in evolutionary terms. I will show that complex emergent individuals such as communities and ecosystems cannot be fully accommodated by conventional population/reproduction models but can be accommodated by variation/persistence models. (shrink)

Following Wallace’s suggestion, Darwin framed his theory using Spencer’s expression “survival of the fittest”. Since then, fitness occupies a significant place in the conventional understanding of Darwinism, even though the explicit meaning of the term ‘fitness’ is rarely stated. In this paper I examine some of the different roles that fitness has played in the development of the theory. Whereas the meaning of fitness was originally understood in ecological terms, it took a statistical turn in terms of reproductive success throughout (...) the 20th Century. This has lead to the ever-increasing importance of sexually reproducing organisms and the populations they compose in evolutionary explanations. I will argue that, moving forward, evolutionary theory should look back at its ecological roots in order to be more inclusive in the type of systems it examines. Many biological systems can only be satisfactorily accounted for by offering a non-reproductive account of fitness. This argument will be made by examining biological systems with very small or transient population structures. I argue this has significant consequences for how we define Darwinism, increasing the significance of survival over that of reproduction. (shrink)

Altruism is one of the most studied topics in theoretical evolutionary biology. The debate surrounding the evolution of altruism has generally focused on the conditions under which altruism can evolve and whether it is better explained by kin selection or multilevel selection. This debate has occupied the forefront of the stage and left behind a number of equally important questions. One of them, which is the subject of this article, is whether the word “selection” in “kin selection” and “multilevel selection” (...) necessarily refers to “evolution by natural selection.” I show, using a simple individual-centered model, that once clear conditions for natural selection and altruism are specified, one can distinguish two kinds of evolution of altruism, only one of which corresponds to the evolution of altruism by natural selection, the other resulting from other evolutionary processes. (shrink)

This article proposes two conditions to assess whether an entity at a level of description is a unit of selection qua interactor. These two conditions make it possible to (1) distinguish biologically relevant entities from arbitrary ones and (2) distinguish units that can _potentially_ enter a selection process from those that have already done so. I show that the classical approaches used in the literature on units and levels of selection do not fare well with respect to either or both (...) of these desiderata. (shrink)

I extend work from Krakauer et al. (2020), who propose a conception of individuality as the capacity to propagate information through time. From this conception, they develop information-theoretic measures. I identify several shortcomings with these measures—in particular, that they are associative rather than causal. I rectify this shortcoming by deriving a causal information-theoretic measure of individuality. I then illustrate how this measure can be implemented and extended in the context of evolutionary transitions in individuality.

Extensive social choice theory is used to study the problem of measuring group fitness in a two-level biological hierarchy. Both fixed and variable group size are considered. Axioms are identified that imply that the group measure satisfies a form of consequentialism in which group fitness only depends on the viabilities and fecundities of the individuals at the lower level in the hierarchy. This kind of consequentialism can take account of the group fitness advantages of germ-soma specialization, which is not possible (...) with an alternative social choice framework proposed by Okasha, but which is an essential feature of the index of group fitness for a multicellular organism introduced by Michod, Viossat, Solari, Hurand, and Nedelcu to analyze the unicellular-multicellular evolutionary transition. The new framework is also used to analyze the fitness decoupling between levels that takes place during an evolutionary transition. (shrink)

A recent development in biology has been the growing acceptance that holobionts, entities comprised of symbiotic microbes and their host organisms, are widespread in nature. There is agreement that holobionts are evolved outcomes, but disagreement on how to characterize the operation of natural selection on them. The aim of this paper is to articulate the contours of the disagreement. I explain how two distinct foundational accounts of the process of natural selection give rise to competing views about evolutionary individuality.

We discuss the meaning of “being alive” and “being dead.” Our primary aim is to pave the way for a sound and accurate ethical debate concerning these two concepts. In particular, we analyze a metabolic approach and a genetic one and discuss the reasons for their failure to constitute a good starting point for successive debates. We argue that any ethical or social discussion of topics involving life and death must introduce cultural constructs such as, on the one hand, the (...) concept of clinical death and, on the other hand, the concept of existence. We argue that these two cultural constructs, although consistent with biomedical knowledge, cannot be ontologically grounded in it. To conclude, we apply our findings to the case of human embryos. (shrink)

According to a shared functionalist view in philosophy of mind, a cognitive system, and cognitive function thereof, is based on the components of the organism it is realized by which, indeed, play a causal role in regulating our cognitive processes. This led philosophers to suggest also that, thus, cognition could be seen as an extended process, whose vehicle can extend not only outside the brain but also beyond bodily boundaries, on different kinds of devices. This is what we call the (...) ‘Externally Extended Cognition Thesis.’ This notion has generated a lively debate. Here, we offer a novel notion of extended cognition, according to which cognition can be seen as being realized outside the brain, but still inside the body. This is what we call the ‘Internally Extended Cognition Thesis’. Not only our thesis but also our approach while defending it is innovative. The argument we offer is supported by recent empirical findings in the life sciences and biomedicine, which suggest that the gut microbiota’s activity has a functional role in regulating our cognitive processes and behaviors. In doing so, we embrace the holobiont-perspective, according to which it is possible to claim that what we call biological individuals are not autonomous entities with clear boundaries, but should rather be seen as networks of multiple interactions among species. Thus, by analyzing different sets of evidence in light of the holobiont-perspective, we argue that the gut microbiota could be seen as a component of our organism. On the basis of the philosophical interpretation of this evidence, however, we also suggest that there are no impediments standing the way of considering the gut microbiota also as a functional extension of our cognitive system. If so, this amounts to extending cognition out of ‘our skull’, though still confining it within ‘our body’: to ‘our gut’. This is an instance of the ‘Internally Extended Cognition Thesis,’ whose benefits for an original theory of extended cognition are discussed. (shrink)

Despite a revived interest in explaining the evolution of anisogamy in recent years (i.e. different—micro and macrogametes), there remain more questions than answers. The topic is important because it is thought to be the foundation of the theory of gender differences and relations. Twelve of these questions are briefly reviewed here—(1) the distinction between sex and sexual types; (2) the distinction between mating types and anisogamy; (3) the possible role of ecological as well as social evolution in proto-gender differences and (...) relations; (4) the life history strategies involved; (5) whether the relevant social relationships are based on conflict or cooperation; (6) the origin and properties of meiosis; (7) the conformity of theories with sex allocation theory; (8) the relevance of multiple levels of selection; (9) appropriate modelling strategies; (10) the relationship between the kind of gametes produced and secondary sexual characteristics; (11) the relevance to humans; and (12) how realistic the search for a single explanation is. It is concluded that polarized sexuality may be a form of trade. (shrink)

This article utilizes three premises. There are commonly ecologically oriented, naturally selected specialized differences in frequency and/or quality as well as sexually selected differences between the sexes. Sex in the sense of coming together and going apart or going apart and coming together is trade in these naturally selected differences, i.e., there is a mating market in sexual species. While such trade is beneficial to the population as a whole, sexual competition and selection is conflict over the profits of that (...) trade and can be detrimental to the population as a whole. These premises yield a naturally selected sex allocation theory of the possible directions and forms of sexual selection, i.e., one that includes costs as well as frequencies. This can explain conventional sex roles, the sex-role reversed, inter- as well as intrasexual selection, and passive as well as active choice. Any of these alternatives may be over mates, over gametes, or both. A hypothetical example based on density dependence relative to resources is provided, one that suggests that centrioles may be a cytoplasmic resource in males in multicellular animals, and which are the target of active choice and the mechanism of manipulation in passive female choice. As a whole, the approach yields a truly general theory of sexual selection, provides an alternative to the mechanism for Fisher’s principle, and gives a theoretical explanation for Mayr’s biological species definition. (shrink)

This paper attempts to reconcile critics and defenders of inclusive fitness by constructing a synthesis that does justice to the insights of both. I argue that criticisms of the regression-based version of Hamilton’s rule, although they undermine its use for predictive purposes, do not undermine its use as an organizing framework for social evolution research. I argue that the assumptions underlying the concept of inclusive fitness, conceived as a causal property of an individual organism, are unlikely to be exactly true (...) in real populations, but they are approximately true given a specific type of weak selection that Hamilton took, on independent grounds, to be responsible for the cumulative assembly of complex adaptation. Finally, I reflect on the uses and limitations of “design thinking” in social evolution research. The debate about the foundations of inclusive fitness theory that has followed in the wake of Nowak, Tarnita and Wilson’s (1) critique has been remarkably polarizing. After several rounds of rebuttals and replies, there is still little evidence of any serious reconciliation between the theory’s critics (2-10) and its defenders (11-24). It doesn’t have to be this way. I believe that, on the main points of disagreement, it is possible to find a way forward that does justice to the insights of both camps. My aim in this paper is to find that way forward. (shrink)

Andrew Bourke’s Principles of Social Evolution identifies three stages that characterize an evolutionary transition in individuality and deploys inclusive fitness theory to explain each stage. The third stage, social group transformation, has hitherto received relatively little attention from inclusive fitness theorists. In this review, I first discuss Bourke’s “virtual dominance” hypothesis for the evolution of the germ line. I then contrast Bourke’s inclusive fitness approach to the major transitions with the multi-level approach developed by Richard Michod, Samir Okasha and others. (...) I suggest that, rather than choosing between these approaches, we should exploit the strengths of both. Finally, I stress the need for a firmer conceptual grasp of the nature of social group transformation. (shrink)

The theories of inclusive fitness and multilevel selection provide alternative perspectives on social evolution. The question of whether these perspectives are of equal generality remains a divisive issue. In an analysis based on the Price equation, Queller argued (by means of a principle he called the separation condition) that the two approaches are subject to the same limitations, arising from their fundamentally quantitative-genetical character. Recently, van Veelen et al. have challenged Queller’s results, using this as the basis for a broader (...) critique of the Price equation, the separation condition, and the very notion of inclusive fitness. Here we show that the van Veelen et al. model, when analyzed in the way Queller intended, confirms rather than refutes his original conclusions. We thereby confirm (i) that Queller’s separation condition remains a legitimate theoretical principle and (ii) that the standard inclusive fitness and multilevel approaches are indeed subject to the same limitations. (shrink)

Various results show the ‘formal equivalence’ of kin and group selectionist methodologies, but this does not preclude there being a real and useful distinction between kin and group selection processes. I distinguish individual- and population-centred approaches to drawing such a distinction, and I proceed to develop the latter. On the account I advance, the differences between kin and group selection are differences of degree in the structural properties of populations. A spatial metaphor provides a useful framework for thinking about these (...) differences: kin and group selection may be conceptualized as large, overlapping regions of K - G space. I then consider some implications of the account, defend it from possible objections, and further argue that the structural features characteristic of both kin and group selection may recur at multiple levels of biological organization. (shrink)

In an incendiary 2010 Nature article, M. A. Nowak, C. E. Tarnita, and E. O. Wilson present a savage critique of the best-known and most widely used framework for the study of social evolution, W. D. Hamilton’s theory of kin selection. More than a hundred biologists have since rallied to the theory’s defence, but Nowak et al. maintain that their arguments ‘stand unrefuted’. Here I consider the most contentious claim Nowak et al. defend: that Hamilton’s rule, the core explanatory principle (...) of kin selection theory, ‘almost never holds’. I first distinguish two versions of Hamilton’s rule in contemporary theory: a special version (HRS) that requires restrictive assumptions, and a general version (HRG) that does not. I then show that Nowak et al. are most charitably construed as arguing that HRS almost never holds, while HRG buys its generality at the expense of explanatory power. While their arguments against HRS are fairly uncontroversial, their arguments against HRG are more contentious, yet these have been largely overlooked in the ensuing furore. I consider the arguments for and against the explanatory value of HRG, with a view to assessing what exactly is at stake in the debate. I suggest that the debate hinges on issues concerning the causal interpretability of regression coefficients, and concerning the explanatory function Hamilton’s rule is intended to serve. (shrink)

In this essay review of Samir Okasha's Agents and Goals in Evolution, I reflect on the rationale for agential thinking in biology, and consider whether the rationale is the same for genes as for organisms. I also discuss Okasha's ingenious examples of the evolution of irrational behaviour, and in particular the evolution of violations of the "independence axiom" of rational choice theory. These examples rely on a crucial distinction between aggregate and idiosyncratic risk.

This paper addresses the foundations of Teleological Individualism, the view that organisms, even non-sentient organisms, are goal-oriented systems while biological collectives, such as ecosystems or conspecific groups, are mere assemblages of organisms. Typical defenses of Teleological Individualism ground the teleological organization of organisms in the workings of natural selection. This paper shows that grounding teleological organization in natural selection is antithetical to Teleological Individualism because such views assume a view about the units of selection on which it is only individual (...) organisms that are units of selection. However, none of the Conventionalist, Reductionist, or Multi-Level Realist theories serve to justify such an assumption. Thus, Teleological Individualism cannot be grounded in natural selection. (shrink)

The Price equation is currently considered one of the fundamental equations —or even the fundamental equation— of evolution. In this article, we explore the role of this equation within cultural evolutionary theory. More specifically, we use it to account for the explanatory power and the theoretical structure of a certain generalised version of dual-inheritance theory. First, we argue that, in spite of not having a definite empirical content, the Price equation offers a suitable formalisation of the processes of cultural evolution, (...) and provides a powerful heuristic device for discovering the actual causes of cultural change and accumulation. Second, we argue that, as a consequence of this, a certain version of the Price equation is the fundamental law of cultural evolutionary theory. In order to support this claim, we sketch the ideal structure of dual-inheritance theory and we stress the unificatory role that the Price equation plays in it. (shrink)

This paper critically supports the modern evolutionary explanation of religion popularised by David Sloan Wilson, by comparing it with those of his predecessors, namely Emile Durkheim and Thomas Hobbes, and to some biological examples which seem analogous to religions as kinds of superorganisms in their own right. The aim of the paper is to draw out a theoretical pedigree in philosophy and sociology that is reflected down the lines of various other evolutionarily minded contributors on the subject of religion. The (...) general theme is of evolved large-scale cooperative structures. A scholarly concern is as follows: Wilson draws on Durkheim, using Calvinism as an example without mentioning Hobbes, but it was Thomas Hobbes who used Calvinism as an example of a leviathanesque religious structure—which is not acknowledged by either Wilson or Durkheim. If there are even any similarities between these authors, there appears to be an omission somewhere which should rightly be accounted for by giving credit to Hobbes where it is due. I issue on conclusion, what it is that makes Wilson’s approach radically different to that it skates on. I also issue it with a cautionary word. (shrink)

Some scholars have raised the potential functional role of fictive kinship for religion, generally. This paper seeks to develop that idea. It is argued in this paper that fictive kinship language in religion (and some other non-religious contexts) recruits traits connected to Hamilton’s rule as it is expressed inHomo sapienspsychology. The effect is that cooperation is augmented within a population that generally shares the same religious worldview. The general position is that if religions are in the business of cooperation and (...) this partially accounts for their evolution and preservation, then it follows that we should take particular note of any significant feature of religions that might lend itself to the cooperation account of religion’s apparent evolutionary success. Fictive kinship is one such feature. (shrink)

Part of Allen et al.’s criticism of Hamilton’s rule makes sense only if we are interested in social adaptation rather than merely social selection. Under the assumption that we are interested in casually modeling social adaptation, I illustrate how graphical causal models of social adaptation can be useful for predicting evolution by adaptation. I then argue for two consequences of this approach given some of the recent philosophical literature. I argue Birch’s claim that the proper way to understand Hamilton’s rule (...) is as providing an organizational framework for causal models is incorrect. I provide an account of a causally adequate decomposition of evolutionary change due to social adaptation and show that my account is superior to Okasha’s. (shrink)

http://dx.doi.org/10.5007/1808-1711.2012v16n1p1 A teoria darwinista tem contribuído para a discussão de problemas nos mais diversos campos filosóficos, entre os quais se inclui a ética e a teoria moral. A sociobiologia e a psicologia evolucionista elucidaram muitos aspectos do comportamento social de diversas espécies animais, a partir de mecanismos como a seleção de parentesco e o altruísmo recíproco que, contudo, são insuficientes para explicar a cooperação no caso humano. Como alternativa, a teoria da dupla herança busca explicar o comportamento humano considerando tanto (...) a biologia quanto as ciências sociais. Segundo esta abordagem, a psicologia social humana é caracterizada por instintos sociais tribais e marcadores simbólicos, que resultam em uma mente que pressupõe princípios morais inatos e universais, selecionados para a vida em grupos orientados por normas sociais, mas que são plasticamente moldados à realidade cultural de cada sociedade. (shrink)

The evolutionary synthesis of the 1930s and 1940s integrated the study of biological microevolution and biological macroevolution into the theoretically consistent and hugely productive field of evolutionary biology. A similar synthesis has yet to occur for the study of culture, and the social sciences remain fragmented and theoretically incompatible. Here, it is suggested that a Darwinian theory of cultural evolution can promote such a synthesis. Earlier non-Darwinian theories of cultural evolution, such as progress theories, lacked key elements of a Darwinian (...) theory of cultural evolution that are necessary to promote a synthesis, while other contemporary theories of cultural evolution, such as memetics, make too stringent neo-Darwinian assumptions that are inconsistent with evidence regarding cultural transmission. Several examples are given which indicate the beginnings of an evolutionary synthesis for culture, where patterns of cultural macroevolution have been explained in terms of underlying cultural microevolutionary forces. Finally, it is argued that experimental simulations of cultural evolution can play an important role in this emerging synthesis. (shrink)

Adaptations can occur at different hierarchical levels (e.g., cells and multicellular organisms), but it can be difficult to identify the level(s) of adaptation in specific cases. A major problem is that selection at a lower level can filter up, creating the illusion of selection at a higher level. We use optimality modeling of the volvocine algae to explore the emergence of genuine group (i.e., colony-level) adaptations. We find that it is helpful to develop an explicit model for what group fitness (...) would be in the absence of group-level relationships between traits and group fitness. We call this “counterfactual fitness,” because in many actual cases of interest there are group-level relationships. Once counterfactual fitness is modeled, the difference between effects that filter up and genuine group selection is explicit and so, therefore, is the distinction between apparent and genuine group adaptations. We call the latter group-specific adaptations. Recognizing group-specific adaptations is important because only group-specific adaptations would cause the lower-level units (cells in this case) to be maladapted if they were to leave the group and enter a global cell-level population. Thus, as group-specific adaptations evolve, they create selective pressure for increased cohesiveness and individuality of groups. This article suggests that group-specific adaptations could be present in the simplest, earliest branching colonial volvocine species, which do not have distinct specialized cells. The article also makes predictions about the kind of empirical evidence needed to support or refute the hypothesis that a particular trait is a group-level adaptation. (shrink)

Why are there so many controversies in evolutionary psychology? Using a couple of concepts from philosophy of science, this paper argues that evolutionary psychology has not reached the stage of mature, normal science, since it does not currently have a unifying research program that guides individual scientists working in the discipline. The argument goes against claims made by certain proponents and opponents of evolutionary psychology, and it is supported by discussion of several examples. The paper notes that just because evolutionary (...) psychology has not reached the stage of normal science, the discipline is nevertheless a source of many progressive theoretical developments and interesting empirical discoveries. (shrink)

A holobiont is a composite organism consisting of a host together with its microbiome, such as a coral with its zooxanthellae. To explain the often intimate integration between hosts and their microbiomes, some investigators contend that selection operates on holobionts as a unit and view the microbiome’s genes as extending the host’s nuclear genome to jointly comprise a hologenome. Because vertical transmission of microbiomes is uncommon, other investigators contend that holobiont selection cannot be effective because a holobiont’s microbiome is an (...) acquired condition rather than an inherited trait. This disagreement invites a simple mathematical model to see how holobiont selection might operate and to assess its plausibility as an evolutionary force. This paper presents two variants of such a model. In one variant, juvenile hosts obtain microbiomes from their parents (vertical transmission). In the other variant, microbiomes of juvenile hosts are assembled from source pools containing the combined microbiomes of all parents (horizontal transmission). According to both variants, holobiont selection indeed causes evolutionary change in holobiont traits. Therefore, holobiont selection is plausibly an effective evolutionary force with either mode of microbiome transmission. The modeling employs two distinct concepts of inheritance, depending on the mode of microbiome transmission: collective inheritance whereby juveniles inherit a sample of the collected genomes from all parents, as contrasted with lineal inheritance whereby juveniles inherit the genomes from only their own parents. A distinction between collective and lineal inheritance also features in theories of multilevel selection. (shrink)

Bourrat and Griffiths :33, 2018) have recently argued that most of the evidence presented by holobiont defenders to support the thesis that holobionts are evolutionary individuals is not to the point and is not even adequate to discriminate multispecies evolutionary individuals from other multispecies assemblages that would not be considered evolutionary individuals by most holobiont defenders. They further argue that an adequate criterion to distinguish the two categories is fitness alignment, presenting the notion of fitness boundedness as a criterion that (...) allows divorcing true multispecies evolutionary individuals from other multispecies assemblages and provides an adequate criterion to single out genuine evolutionary multispecies assemblages. A consequence of their criterion is that holobionts, as conventionally defined by hologenome defenders, are not evolutionary individuals except in very rare cases, and for very specific host-symbiont associations. This paper is a critical response to Bourrat and Griffiths’ arguments and a defence of the arguments presented by holobiont defenders. Drawing upon the case of the hologenomic basis of the evolution of sanguivory in vampire bats, I argue that Bourrat and Griffiths overlook some aspects of the biological nature of the microbiome that justifies the thesis that holobionts are evolutionarily different to other multispecies assemblages. I argue that the hologenome theory of evolution should not define the hologenome as a collection of genomes, but as the sum of the host genome plus some traits of the microbiome which together constitute an evolutionary individual, a conception I refer to as the stability of traits conception of the hologenome. Based on that conception I argue that the evidence presented by holobiont defenders is to the point, and supports the thesis that holobionts are evolutionary individuals. In this sense, the paper offers an account of the holobiont that aims to foster a dialogue between hologenome advocates and hologenome critics. (shrink)

The units or levels of selection debate concerns the question of what kind of biological systems are stable enough that part of their evolution is a result of the process of natural selection acting at their level. Traditionally, the debate has concerned at least two different, though related, questions: the question of the level at which interaction with the environment occurs, and the question of the level at which reproduction occurs. In recent years, biologists and philosophers have discussed a new (...) aspect of this debate, namely the possibility that certain multi-species consortia formed by a host and its microbiome may act as a unit of selection. This thesis, however, has not been without criticism, as it is doubtful that such consortia could meet the conditions required to achieve the degree of stability that would allow them to experience natural selection. The purpose of this paper is to systematically examine such criticisms and to defend the thesis that the holobiont/hologenome can act as a genuine level of selection both in the form of an interactor and in the form of a reproducer. To do so, it will be argued that the microbiome should be characterized in functional rather than taxonomic terms. (shrink)

The hologenome concept of evolution is a hypothesis about the evolution of animals and plants. It asserts that the evolution of animals and plants was partially triggered by their interactions with their symbiotic microbiomes. In that vein, the hologenome concept posits that the holobiont (animal host + symbionts of the microbiome) is a unit of selection. -/- The hologenome concept has been severely criticized on the basis that selection on holobionts would only be possible if there were a tight transgenerational (...) host-genotype-to-symbiont-genotype connection. As our current evidence suggests that this is not the case for most of the symbiont species that compose the microbiome of animals and plants, the opportunity for holobiont selection is very low in relation to the opportunity for selection on each of the species that compose the host microbiome. Therefore, holobiont selection will always be disrupted ‘from below’, by selection on each of the species that compose the microbiome. -/- This thesis constitutes a conceptual effort to defend philosophically the hologenome concept. I argue that the criticism according to which holobiont selection requires tight transgenerational host-genotype-to-symbiont-genotype connection is grounded on a metaphysical view of the world according to which the biological hierarchy needs to be nested, such that each new level of selection includes every entity from below. Applied to hologenomes, it entails that the hologenome is a collection of genomes, and selection of hologenomes is assumed to entail cospeciation of the host with the species that constitute its microbiome. -/- Against that interpretation, I propose the ‘stability of traits’ account, according to which hologenome evolution is the result of the action of natural selection in a non-nested hierarchical world. In that vein, hologenome evolution does not entail cospeciation, and thus it does not require tight transgenerational host-genotype-to-symbiont-genotype connection. By embracing a multilevel selection perspective, I argue that hologenome evolution results from the simultaneous action of natural selection on each of the lineages that compose the microbiome, and on the assemblage composed by the host genome plus the functional traits of its microbiome. Hologenome selection occurs when the evolution of the traits of the microbiome result from their effects on the fitness of the host, and it can take the form of multilevel selection 1, or multilevel selection 2. In both cases, hologenome selection entails the evolution of microbiome traits, as well as evolution of the host genome, rather than cospeciation of lineages. (shrink)

Contemporary biological research has suggested that some host–microbiome multispecies systems (referred to as “holobionts”) can in certain circumstances evolve as unique biological individual, thus being a unit of selection in evolution. If this is so, then it is arguably the case that some biological adaptations have evolved at the level of the multispecies system, what we call hologenomic adaptations. However, no research has yet been devoted to investigating their nature, or how these adaptations can be distinguished from adaptations at the (...) species-level (genomic adaptations). In this paper, we cover this gap by investigating the nature of hologenomic adaptations. By drawing on the case of the evolution of sanguivory diet in vampire bats, we argue that a trait constitutes a hologenomic adaptation when its evolution can only be explained if the holobiont is considered the biological individual that manifests this adaptation, while the bacterial taxa that bear the trait are only opportunistic beneficiaries of it. We then use the philosophical notions of emergence and inter-identity to explain the nature of this form of individuality and argue why it is special of holobionts. Overall, our paper illustrates how the use of philosophical concepts can illuminate scientific discussions, in the trend of what has recently been called metaphysics of biology. (shrink)

Reduction and reductionism have been central philosophical topics in analytic philosophy of science for more than six decades. Together they encompass a diversity of issues from metaphysics and epistemology. This article provides an introduction to the topic that illuminates how contemporary epistemological discussions took their shape historically and limns the contours of concrete cases of reduction in specific natural sciences. The unity of science and the impulse to accomplish compositional reduction in accord with a layer-cake vision of the sciences, the (...) seminal contributions of Ernest Nagel on theory reduction and how they strongly conditioned subsequent philosophical discussions, and the detailed issues pertaining to different accounts of reduction that arise in both physical and biological science (e.g., limit-case and part-whole reduction in physics, the difference-making principle in genetics, and mechanisms in molecular biology) are explored. The conclusion argues that the epistemological heterogeneity and patchwork organization of the natural sciences encourages a pluralist stance about reduction. (shrink)

It is an unfortunate fact of academic life that there is a sharp divide between science and philosophy, with scientists often being openly dismissive of philosophy, and philosophers being equally contemptuous of the naivete ́ of scientists when it comes to the philosophical underpinnings of their own discipline. In this paper I explore the possibility of reducing the distance between the two sides by introducing science students to some interesting philosophical aspects of research in evolutionary biology, using biological theories of (...) the origin of religion as an example. I show that philosophy is both a discipline in its own right as well as one that has interesting implications for the understanding and practice of science. While the goal is certainly not to turn science students into philoso- phers, the idea is that both disciplines cannot but benefit from a mutual dialogue that starts as soon as possible, in the classroom. (shrink)

The propensity interpretation of fitness (PIF) is commonly taken to be subject to a set of simple counterexamples. We argue that three of the most important of these are not counterexamples to the PIF itself, but only to the traditional mathematical model of this propensity: fitness as expected number of offspring. They fail to demonstrate that a new mathematical model of the PIF could not succeed where this older model fails. We then propose a new formalization of the PIF that (...) avoids these (and other) counterexamples. By producing a counterexample-free model of the PIF, we call into question one of the primary motivations for adopting the statisticalist interpretation of fitness. In addition, this new model has the benefit of being more closely allied with contemporary mathematical biology than the traditional model of the PIF. (shrink)

Even after Willard Quine’s critique of the analytic-synthetic distinction in “Two Dogmas of Empiricism,” Wilfrid Sellars maintained some forms of analyticity or truth in virtue of meaning. In this article, I aim to reconstruct (a) his neglected account of the analytic-synthetic distinction and the revisability of analytic sentences, (b) its connection to his inferentialist account of meaning, and (c) his response to Quine. While Sellars’s account of the revisability of analytic sentences bears certain similarities to Carnap’s and Grice and Strawson’s (...) accounts, it is still striking in that it is part of his broader picture of how our language develops dynamically in our ongoing inquiry. I aim to show how it relates to his account of how the diachronic continuity in meaning, language, or conceptual framework can be preserved through a revision of analytic sentences and to his evolutionary account of the development of our language or conceptual framework. (shrink)

Since, it has become clear that individuality is not to be considered as a given, but rather as something which needs to be explained. How has individuality emerged through evolution, and how has it subsequently been maintained? In particular, why is it that multicellular organisms appeared and persisted, despite the obvious interest of each cell of favoring its own replication? Several biologists see the immune system as one of the key components for explaining the maintenance of multicellular organisms’ individuality. Indeed, (...) the immune system exerts a constant surveillance on all the constituents of the organism, including “cheaters” like tumor cells, which favor their own replication at the expense of the whole organism. In most cases, the immune system eliminates those cheaters. This is the “immune surveillance” hypothesis, first suggested by Burnet and Thomas. In this paper, I account for recent findings on immune surveillance in order to determine the precise role of the immune system in the emergence and maintenance of individuality. This investigation gives rise to a critical question for the domain of biological individuality: should immunity be considered as something unique to multicellular organisms? If it is indeed unique, should the multicellular organism be considered as an individual to a higher degree than any other living entity? If it is not unique, what precisely are the equivalents of the immune system in other living individuals? (shrink)

A crucial question for a process view of life is how to identify a process and how to follow it through time. The genidentity view can contribute decisively to this project. It says that the identity through time of an entity X is given by a well-identified series of continuous states of affairs. Genidentity helps address the problem of diachronic identity in the living world. This chapter describes the centrality of the concept of genidentity for David Hull and proposes an (...) extension of Hull’s view to the ubiquitous phenomenon of symbiosis. Finally, using immunology as a key example, it shows that the genidentity view suggests that the main interest of a process approach is epistemological rather than ontological and that its principal claim is one of priority, namely that processes precede and define things, and not vice versa. (shrink)

The traditional practice of establishing morphological types and investigating morphological organization has found new support from evolutionary developmental biology (evo-devo), especially with respect to the notion of body plans. Despite recurring claims that typology is at odds with evolutionary thinking, evo-devo offers mechanistic explanations of the evolutionary origin, transformation, and evolvability of morphological organization. In parallel, philosophers have developed non-essentialist conceptions of natural kinds that permit kinds to exhibit variation and undergo change. This not only facilitates a construal of species (...) and higher taxa as natural kinds, but also broadens our perspective on the diversity of kinds found in biology. There are many different natural kinds relevant to the investigative and explanatory aims of evo-devo, including homologues and developmental modules. (shrink)

There is an active research program currently underway, which treats cancer progression as an evolutionary process. This contribution investigates the ways that cancer progression is like and unlike evolution in other contexts. The aim is to take a multi-level perspective on cancer, investigating the levels at which selection may be acting, the unit or target of selection, the relative roles of selection and drift, and the idea that cancer progression may be a by-product of selection at other levels of organization.

The theory of evolution settled at what was thought to be its definitive form after the affiliation of Darwin’s theory with the new science of genetics. This historical event explains not only the success but also the vulnerability of evolutionary theory. The close affinity with genetics helped to provide the tools required for managing phylogenetic evolution, which was controlled by the molecular machinery of the genome, localized in most cells of each individual. This setup worked well for organizing the basics (...) of Darwinism but it faltered when applied to the evolution of culture. This is so because, as I argue in this article, under the given conditions the active support of a system of similar complexity as the genome would be required. At some point in the process of evolution the large brains of primates may have begun experimenting with phenotypic equivalents of genomic functions like memory, adaptation, fitness, processing information, and organizing replication. In this way the brain may have weakened its ties to the genome and by exploiting its new status have become, so to speak, for the phenotype what the genome is for the genotype, thus preparing the stage on which the complexity of human behavior might unfold. (shrink)

Evolutionary biology is a field currently animated by much discussion concerning its conceptual foundations. On the one hand, we have supporters of a classical view of evolutionary theory, whose backbone is provided by population genetics and the so-called Modern Synthesis (MS). On the other hand, a number of researchers are calling for an Extended Synthe- sis (ES) that takes seriously both the limitations of the MS (such as its inability to incorporate developmental biology) and recent empirical and theoretical research on (...) issues such as evolvability, modularity, and self-organization. In this article, I engage in an in-depth commentary of an influential paper by population geneticist Michael Lynch, which I take to be the best defense of the MS-population genetics position published so far. I show why I think that Lynch’s arguments are wanting and propose a modification of evolutionary theory that retains but greatly expands on population genetics. (shrink)